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Novel fungi from an ancient niche: Cercosporoid and related sexual morphs on ferns

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The fern flora of the world (Pteridophyta) has direct evolutionary links with the earliest vascular plants that appeared in the late Devonian. Knowing the mycobiota associated to this group of plants is critical for a full understanding of the Fungi. Nevertheless, perhaps because of the minor economic significance of ferns, this niche remains relatively neglected by mycologists. Cercosporoid fungi represent a large assemblage of fungi belonging to the Mycosphaerellaceae and Teratosphaeriaceae (Ascomycota) having cercospora-like asexual morphs. They are well-known pathogens of many important crops, occurring on a wide host range. Here, the results of a taxonomic study of cercosporoid fungi collected on ferns in Brazil are presented. Specimens were obtained from most Brazilian regions and collected over a 7-yr period (2009–2015). Forty-three isolates of cercosporoid and mycosphaerella-like species, collected from 18 host species, representing 201 localities, were studied. This resulted in a total of 21 frond-spotting taxa, which were identified based on morphology, ecology and sequence data of five genomic loci (actin, calmodulin, ITS, LSU and partial translation elongation factor 1-α). One novel genus (Clypeosphaerella) and 15 novel species (Cercospora samambaiae, Clypeosphaerella sticheri, Neoceratosperma alsophilae, N. cyatheae, Paramycosphaerella blechni, Pa. cyatheae, Pa. dicranopteridis-flexuosae, Pa. sticheri, Phaeophleospora pteridivora, Pseudocercospora brackenicola, Ps. paranaensis, Ps. serpocaulonicola, Ps. trichogena, Xenomycosphaerella diplazii and Zasmidium cyatheae) are introduced. Furthermore, 11 new combinations (Clypeosphaerella quasiparkii, Neoceratosperma yunnanensis, Paramycosphaerella aerohyalinosporum, Pa. dicranopteridis, Pa. gleicheniae, Pa. irregularis, Pa. madeirensis, Pa. nabiacense, Pa. parkii, Pa. pseudomarksii and Pa. vietnamensis) are proposed. Finally, nine new host associations are recorded for the following known fungal species: Cercospora coniogrammes, Cercospora sp. Q, Ps. abacopteridicola, Ps. lygodiicola and Ps. thelypteridis.
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... based on a multi-locus DNA sequence dataset of five genomic loci (ITS, actA, cmdA, his3 and tef1) of a large sampling of species. Since then, numerous molecular examinations of Cercospora species have been carried out based on multi-gene approaches (Dianese et al. 2014, Bakhshi et al. 2015a, Shivas et al. 2015, Soares et al. 2015, Albu et al. 2016, Guatimosim et al. 2016, Nguanhom et al. 2016, Guillin et al. 2017. Bakhshi et al. (2018) applied three more potential candidate gene regions including gapdh, rpb2, and tub2 to perform an eight-gene phylogeny (ITS,actA,cmdA,gapdh,his3,rpb2,tef1 and tub2) for Cercospora species. ...
... References: Ellis 1971, Chupp 1954, Crous & Braun 2003 Groenewald et al. 2013, Bakhshi et al. 2015a, Guatimosim et al. 2016, Nguanhom et al. 2016 Leaf spots amphigenous, circular, 3-6 mm, grey with definite border and yellow halo. Mycelium internal. ...
... (Videira et al. 2017), showed Pseudocercospora nephrolepidicola warranted transfer to a new genus, Pteridopassalora. Many species of Passalora and Pseudocercospora that have been reported on ferns in recent years (Crous et al. 2010, Kirschner & Liu 2014, Kirschner & Wang 2015, Guatimosim et al. 2016, have morphological characteristics that resembling those of Pteridopassalora. Further studies are required to clarify their molecular phylogeny. ...
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... In this study, a transverse leaf section for the first time in Virosphaerella revealed the presence of hyphae within the intercellular spaces of the host tissues, as also recorded in some other Mycospharellaceae (Kirschner & Okuda 2013). The morphological characteristics were similar to other teleomorphic Mycosphaerellalike ascomycetes such as Clypeosphaerella sticheri Guatim., R.W. Barreto & Crous occurring on members of other fern genera within the Gleicheniaceae (Guatimosim et al. 2016), but the phylogenetic hypotheses (Figs. 3, 4) indicate that these other fungi and our specimens represent different species in different lineages. In each of these diverse mycosphaerellaceous genera with species on Gleichenicaceae, there are also other species associated with seed plants. ...
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